Notes on the position of the true freshwater crabs within the brachyrhynchan Eubrachyura (Crustacea: Decapoda: Brachyura)

Cladistic and phenetic relationships of 51 eubrachyuran crab genera, comprising 36 genera of marine crabs and 18 genera of true freshwater crabs from 7 families, were investigated using 121 parsimony-informative adult morphological characters. The data matrix was subjected to four different treatments: (1) a cladistic analysis with a combination of unordered and ordered characters, (2) a cladistic analysis with all characters unordered, (3) neighbour-joining, and (4) UPGMA phenetic analyses. The parsimony analysis conducted with a combination of ordered and unordered characters produced a set of hypotheses which supported monophyly of a Pseudothelphusidae+Potamoidea clade. Furthermore, exemplar genera of the Bythograeidae and Pinnotheridae formed an unresolved polytomy with the Pseudothelphusidae+Potamoidea group, the Thoracotremata. The trichodactylid freshwater crabs were positioned as the sister taxon of the basal portunoid Carcinus, but were unresolved relative to other portunoids and geryonids. Second, the parsimony analysis conducted with all characters unordered resulted in a [bythograeid, pseudothelphusid+potamoid, pinnotherid, thoracotreme] group with no hierarchical resolution, which in turn formed a polytomy with a goneplacid+portunoid clade and a polyphyletic Xanthoidea. And third, phenetic groupings of the eubrachyuran genera invariably placed the pseudothelphusids with the potamoids, and this clustered with a group containing the thoracotremes (either in whole or part). Support was thus found for morphological connections among the nontrichodactylid freshwater crabs, thoracotremes, bythograeids, and pinnotherids, and for the placement of the trichodactylids within the Portunoidea. These two latter findings (that used a range of genera from each family) are broadly congruent with a previous cladistic analysis of selected eubrachyuran familial groundpatterns that used a basal exemplar of each marine and freshwater crab family (Sternberg et al., 1999). However, it is clear that the large scale homoplasy identified here may nullify any reliable hypothesis of brachyrhynchan groupings at this stage.     

Global diversity of crabs (Crustacea: Decapoda: Brachyura) in freshwater

An assessment of the global freshwater crab diversity is presented. A total of 1,476 species in 14 families are currently known from all zoogeographical regions (except Antarctica), including 1,306 species in eight exclusively freshwater families (Pseudothelphusidae, Trichodactylidae, Potamonautidae, Deckeniidae, Platythelphusidae, Potamidae, Gecarcinucidae and Parathelphusidae). Estimates of true freshwater crab diversity including likely numbers of undescribed taxa suggest that the field remains largely in a “discovery” phase. Main ideas on the origins, diversification, and phylogeny of true freshwater crabs are briefly discussed. The economic importance of freshwater crabs is also highlighted. Guest editors: E. V. Balian, C. Lévêque, H. Segers & K. Martens Freshwater Animal Diversity Assessment     

A conservation assessment of the freshwater crabs of southern Africa (Brachyura: Potamonautidae)

Recent taxonomic revisions of the freshwater crabs of southern Africa (Angola, Botswana, Lesotho, Mozambique, Namibia, South Africa, Swaziland, Zambia and Zimbabwe) allow accurate depictions of their diversity, distribution patterns and conservation status. The southern African region is home to nineteen species of freshwater crabs all belonging to the genus Potamonautes (family Potamonautidae). These crabs show high levels of species endemism (84%) to the southern African region and to the country of South Africa (74%). The conservation status of each species is assessed using the IUCN (2003) Red List criteria, based on detailed compilations of the majority of known specimens. The results indicate that one species should be considered vulnerable, fifteen species least concern and three species data deficient. The results have been utilized by the IUCN for Red Lists, and may prove useful when developing a conservation strategy for southern Africa’s endemic freshwater crab fauna.     

Phylogenetic analysis of the Mexican freshwater crabs of the tribe Pseudothelphusini (Decapoda: Brachyura: Pseudothelphusidae)

The Mexican pseudothelphusid crabs are classified in one subfamily, three tribes, and 13 genera. Up to now, 56 species have been recognized, distributed in a strictly Neotropical pattern, with some of them reaching the state of Sonora on the western slope of Mexico. The tribe Pseudothelphusini is the most diverse, with five genera and 35 species, all of them endemic to Mexico: the two most species‐rich genera are Pseudothelphusa, with 23 species, and Tehuana, with eight species; Epithelphusa includes two species, whereas Disparithelphusa and Smalleyus are monotypic. The Pseudothelphusini lack an updated systematic revision, which could serve as a framework to analyse the monophyletic origin of the group, to clarify the relationships among genera and species, as well as to resolve the taxonomic status of various species complexes. In the present study, an exhaustive morphological revision was conducted using somatic and sexual characters. A phylogenetic analysis was performed using 77 characters and 183 character states, taken from 41 species. Ten trees of the same length were obtained using PAUP 4.0 through a heuristic search. The results show that the tribe as it is actually known constitutes a paraphyletic group, in which the species of Epithelphusa and Pseudothelphusa puntarenas are excluded from the internal group. According to the obtained results, the tribe Pseudothelphusini s.s. includes five genera: Smalleyus, Pseudothelphusa, Tehuana, and two new ones to accommodate Pseudothelphusa galloi and Pseudothelphusa sulcifrons, respectively. This new arrangement considers the provisional suppression of the genus Disparithelphusa, which remained as another species of Pseudothelphusa throughout the cladistic analysis. The phylogenetic results show a strong congruence with the distribution of the species, in several cases grouping species that form morphological clines along a geographical gradient. The previously proposed southern origin of the tribe Pseudothelphusini gains support with the results obtained. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010, 160 , 457–481     

Ultrastructure of spermatozoa and spermatophores of old world freshwater crabs (Brachyura: Potamoidea: Gecarcinucidae,Potamidae, and Potamonautidae)

We investigated the ultrastructure of spermatozoa and spermatophores of 19 palaeotropical freshwater crab species [12 species of the Gecarcinucidae, 6 of the Potamidae (Potamiscinae), and 1 species of the Potamonautidae (Deckeniinae: Hydrothelphusini)]. The investigated Potamiscinae have densely packed coenospermic spermatophores with the exception of Thaiphusa sirikit and Johora singaporensis that exhibit cleistospermia. In contrast, in the Gecarcinucidae the spermatozoa are loosely embedded in a mucous matrix. The gecarcinucid and potamiscine sperm differ, furthermore, in acrosomal structure and size. The acrosome in the Gecarcinucidae is much smaller and spherical, while the larger acrosome in the Potamiscinae has the tendency to be depressed. In the Potamiscinae, an additional middle acrosomal zone evolved between the acrosome ray zone and the outer acrosomal zone. Within the Gecarcinucidae, a differentiation into two groups (Gecarcinucinae and Parathelphusinae) is not supported by the present spermatological data. The sperm morphology of Hydrothelphusa aff. madagascariensis (Potamonautidae: Deckeniinae) differs from Potamonautes sidneyi (Potamonautidae: Potamonautinae) in acrosomal size and shape, and in the absence of a periopercular rim. A closer relationship of Deckeniinae and Gecarcinucidae cannot be confirmed by spermatology. J. Morphol., 2009. © 2008 Wiley‐Liss, Inc.     

Phylogenetics of Cancer crabs (Crustacea: Decapoda: Brachyura).

We used morphological, mitochondrial DNA sequence, paleontological, and biogeographical information to examine the evolutionary history of crabs of the genus Cancer. Phylogenies inferred from adult morphology and DNA sequence of the cytochrome oxidase I (COI) gene were each well resolved and well supported, but differed substantially in topology. Four lines of evidence suggested that the COI data set accurately reflected Cancer phylogeny: (1) in the phylogeny inferred from morphological data, each Atlantic species was sister taxon to an ecologically similar Pacific species, suggesting convergence in morphology; (2) a single trans-Arctic dispersal event, as indicated by the phylogeny inferred from COI, is more parsimonious than two such dispersal events, as inferred from morphology; (3) test and application of a maximum likelihood molecular clock to the COI data yielded estimates of origin and speciation times that fit well with the fossil record; and (4) the tree inferred from the combined COI and morphology data was closely similar to the trees inferred from COI, although notably less well supported by the bootstrap. The phylogeny inferred from maximum likelihood analysis of COI suggested that Cancer originated in the North Pacific in the early Miocene, that the Atlantic species arose from a North Pacific ancestor, and that Cancer crabs invaded the Atlantic from the North Pacific 6-12 mya. This inferred invasion time is notably prior to most estimates of the date of submergence of the Bering Strait and the trans-Arctic interchange, but it agrees with fossil evidence placing at least one Cancer species in the Atlantic about 8 mya.     

The freshwater palaemonid prawns (Crustacea: Decapoda: Caridea) of Myanmar

The taxonomy of the palaemonid freshwater prawns of Myanmar is reviewed and 19 species are recorded. These belong to three genera, with 17 species of Macrobrachium, one species of Palaemonetes and one species of Leptocarpus. The identities of Palaemon spinipes birmanicus Schenkel, 1902, and Palaemon naso Kemp, 1918, are clarified with both recognised as distinct taxa. The taxonomy of several poorly known species of Macrobrachium,viz. M. villosimanus (Tiwari, 1949), M. peguense (Tiwari, 1952), M. hendersoni (De Man, 1906) and M. platyrostrum (Tiwari, 1952) is revised and discussed. Five species are reported from Myanmar for the first time, viz. Macrobrachium lamarrei (H. Milne-Edwards, 1837), M. dayanum (Henderson, 1893), M. lanchesteri (De Man, 1911), M. nipponense (De Haan, 1894), and Palaemonetes sinensis (Sollaud, 1911). One new species, Macrobrachium lanatum, which is closely allied to M. scabriculum (Heller, 1862) and M. dolichodactylus (Hilgendorf, 1879), is described and figured.     

Molecular and morphometric data demonstrate the presence of cryptic lineages among freshwater crabs (Decapoda: Potamonautidae: Potamonautes) from the Drakensberg Mountains, South Africa

The evolutionary importance of cryptic taxa is well documented, yet few studies have examined them with a view to conservation. In the present study, the significance of cryptic speciation in freshwater crabs is examined. Allozyme and 16S rRNA sequence data were used to explore the degree of population differentiation between mountain stream populations of two distinct freshwater crab species. Marked patterns of differentiation were evident among populations; those in close geographical proximity were characterized by the fixation of alternate alleles at certain loci, indicating that currently there is no gene flow among populations. Both allozyme and sequence data provide evidence for the recognition of at least five distinct evolutionary lineages with pronounced levels of genetic differentiation. Morphometric data failed to detect any distinct geographically intermediate population groupings. Our findings indicate the presence of five phylogeographical units, all worthy of conservation, three of which are evolutionarily significant.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 78 , 129–147.     

Evolution of sperm morphology in potamid freshwater crabs (Crustacea: Brachyura: Potamoidea)

We investigated sperm cells and spermatophores of four species of Old World freshwater crabs belonging to three different genera of the subfamily Potaminae (family Potamidae). Characters previously believed to be apomorphic for the potamid subfamily Potamiscinae were also found to occur in the Potaminae. To infer the morphological ancestral character state combination of the Potamidae, ancestral character state analysis of four different sperm traits was performed, based on a 16S rDNA phylogeny of the investigated species. Comparing molecular phylogeny and character state distribution, several cases of convergent evolution could be identified. The densely packed, coenospermic spermatophores and the occurrence of a ‘tongue‐and‐groove’ connection between operculum and acrosomal zones are probably apomorphies for the whole Potamidae. The spermatozoa of Socotrapotamon socotrense show several unique characters. We also analysed the evolution of acrosome size. The sperm cells of the Potamidae and their sister‐group Gecarcinucidae only slightly overlap in acrosome size. Within the investigated species, the ‘East Asia’ subclade (subfamily Potamiscinae) developed significantly larger acrosomes than the subfamily Potaminae. Our results suggest that the use of brachyuran acrosome morphology for phylogenetic inference at the family level is strongly affected by small sample size, and by convergent character evolution. © 2010 The Linnean Society of London, Zoological Journal of the Linnean Society, 2010.     

The collision of the Indian plate with Asia: molecular evidence for its impact on the phylogeny of freshwater crabs (Brachyura: Potamidae)

Aim We used molecular data to answer the following questions: (1) Is morphology‐based (and to some extent, geography‐based) classification of the freshwater crab family Potamidae congruent with a molecular phylogeny? (2) What historical biogeographical event could have shaped this phylogeny? Location Material from the entire geographical range of the family Potamidae was analysed, including specimens from East Asia (China, Taiwan, the Ryukyus), Southeast Asia, South Asia (northern India, the Middle East and Near East), North Africa, and southern Europe. Methods Mitochondrial DNA sequences encoding 503 base pairs (excluding the variable regions) of the large subunit rRNA (16S rRNA) gene were obtained from 72 species belonging to 49 potamid genera, representing 51% of all known genera in this species‐rich family. Sequences were compared by means of phylogenetic analyses (minimum evolution, Bayesian inference, maximum likelihood and maximum parsimony) and Bayesian relaxed molecular clock estimates. Results The family Potamidae was found to be monophyletic with two major lineages, and there was support for the recognition of two mostly allopatric subfamilies, Potaminae and Potamiscinae. This is largely consistent with the current classification proposed. The ‘Potamiscinae’ clade comprised three subclades: (1) a well‐supported ‘eastern Asia’ subclade that included species from the eastern part of the range (China, Taiwan, the Ryukyus, the Philippines, Indochina, Malay Peninsula, northern India and Myanmar/Burma); (2) a weakly supported ‘Sunda Shelf islands’ subclade that included species from the larger Southeast Asian islands on the Sunda Shelf (Borneo, Sumatra and Java); and (3) a ‘Socotra’ subclade that comprised only Socotrapotamon from Socotra Island, off the north‐east coast of Africa. Main conclusions The discrete distribution of the two subfamilies in Europe/Asia is hypothesized to be the result of vicariance due to the collision of the Indian tectonic plate with the Asian continent, and the orogeny that caused the separation of the two freshwater crab lineages around 22.8 Ma. Within the Potamiscinae, the ‘Sunda Shelf islands’ subclade separated from other potamiscines around 21.1 Ma; and the endemic fauna of the East Asian islands (Taiwan, the Ryukyus and mainland Japan) was isolated from the Asian continent c. 8.4 Ma, following the opening of the Okinawa Trough. The ‘Socotra’ subclade diverged from the ‘eastern Asia’ subclade at 19.1 Ma during the Miocene. Its taxonomic position, however, remains unclear as the members of this clade possess the key potamine character of a transverse ridge on thoracic sternite 8, suggesting that this may in fact be a relict potamid group.     

Mitochondrial sequence data reveals the phylogeny of the Asian Helice group of crabs (Decapoda: Brachyura: Varunidae)

The taxonomy of the Helice s. l. crabs has been recently revised, subdividing the Asian species in three genera, Helice , Helicana and Pseudohelice . The last one is widespread throughout the Indo-West Pacific Ocean and a single species, Pseudohelice subquadrata , is found in Asia. The other two genera appear to be endemic to Asia. However, phylogenetic affinities among the Asian representatives of Helice s. l. are poorly understood, and different morphology-based hypotheses of relationships remain untested. In this study, we used sequence data from a mitochondrial gene-block spanning three different genes [16S rDNA- trnL ₁ - IGS -nad1 ] to perform phylogenetic analyses. The results provide independent molecular evidence supporting the splitting of the Asian species into Helice and Helicana . Helice is suggested to be paraphyletic with respect to P. subquadrata . In addition, the heterogeneity of the genus Pseudohelice is shown when including the East Asian and East African counterparts. We propose that the two genera, Helice and Pseudohelice , are not monophyletic in their current composition.     

Not only for males: the pleon‐holding mechanism in female calappids (Crustacea: Decapoda: Brachyura)

Adult female Calappidae were examined with light microscopy and microcomputed tomography for the presence of a pleon‐holding mechanism. A typical press‐button consisting of a protrusion on the fifth thoracic sternite and a socket on the sixth pleon segment was found in species of the genera Calappa, Calappula, Cycloes, Mursia, and Paracyclois. Within the genus Calappa, 15 of 17 species showed a complete press‐button mechanism, whereas in two species only a sternal button or none of the holding structures could be observed. The functionality of the mechanism was assessed by manually flexing and opening the pleon. This revealed a lack of function in species with fully developed holding structures, while others exhibited full function. Additionally, signs of abrasive wear could be identified with scanning electron microscopy on the surface of the holding structures of Calappa hepatica. The documentation of pleon‐holding structures of varying degrees of functionality in adult female eubrachyurans provides new insight into a morphological characteristic known exclusively from Brachyura.     

Phylogénie des touracos (Aves, Musophagidae). Analyse des caractères morphologiques

Phylogeny of turacos (Aves, Musophagidae). Analysis of morphological characters
A cladistic analysis of the 23 species of turacos (Musophagidae) using 34 morphological characters produces, after successive weighting, 4 shortest-length trees (consistency index = 0.76) that present two major clades: the first one with 'grey' turacos (genus Crinifer, Corythaixoides ), the other one with the great blue turaco (genus Corythaeola ) and the turacine bearers turacos (genus Tauraco , Musophaga , Ruwenzorornis , Gallirex ). This result also suggests that the species ' persa ' is not valid; T. schalowi , T. livingstonii , T. corythaix which were included in this species are more closely related to T. fisheri and T. schuettii than they are to T. persa . The polymorphism of these green turacos is also discussed. Ruwenzorornis johnstoni and Gallirex porphyreolophus do not belong to the Tauraco clade nor to the Musophaga clade, but they are closer to the first than to the second. The species Corythaixoides leucogaster is closer to the Crinifer species than to the other Corythaixoides species.     

New species and new records of pinnotherid crabs (Crustacea: Decapoda: Brachyura) from the Yellow Sea

Five species belonging to two genera of pinnotherid crabs are reported from the Yellow Sea. Two are new to science, namely: Sakaina glabra sp. nov. and Sakaina granulata sp. nov.; while three species are recorded for the first time from China: Pinnaxodes majorOrtmann, 1894, Pinnaxodes mutuensisSakai, 1939, and Sakaina japonicaSerène, 1964. Sakaina glabra can be easily distinguished from congeners by lacking a thick rim of pubescence along the anterolateral margin of the carapace. Sakaina granulata closely resembles S. asiatica (Sakai, 1933), but differs mainly in having a row of granule-like denticles along the anterolateral margin of its carapace.     

Female reproductive morphology of coral‐inhabiting gall crabs (Crustacea: Decapoda: Brachyura: Cryptochiridae)

Gall crabs are obligate associates of stony corals in which they induce skeletal modifications. In some cryptochirid species, females live in open depressions accessible to males; while in others, females are rather isolated in semiclosed galls, which necessitates elaborate sperm storage capabilities by the female. In this study we investigate the female gross morphology and reproductive systems of Fungicola syzygia lodged in semiclosed flattened pits, Opecarcinus cathyae with semiopen pits and Pseudocryptochirus viridis from shallow open depressions using line drawings and histological methods. The general morphology of the cryptochirids' reproductive systems is uniform and conforms to other thoracotreme brachyurans: paired muscular vaginae of the concave pattern lead from the sternal gonopores into paired seminal receptacles where sperm is stored. The seminal receptacle is internally lined by distinct types of epithelia: a cuticle underlined by a columnar epithelium ventrally, a monolayered secretory epithelium dorsally and a multilayered transfer tissue where the oviducts enter the seminal receptacle. In all studied specimens, the seminal receptacle contained free spermatozoa; however, in specimens of Pseudocryptochirus viridis it also contained spermatophores, indicating a recent insemination. In contrast to most other brachyurans ovaries of the investigated cryptochirids extend into the pleon. The specific degree of ovary extension differs between the studied species and is closely related to female body shape.